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LET LET12382.Pdf (13.79Mb) After 100 years: a detailed view of an eumalacostracan crustacean from the Upper Jurassic Solnhofen Lagerstätte with raptorial appendages unique to Euarthropoda PAULA G. PAZINATO , CLÉMENT JAUVION , GÜNTER SCHWEIGERT, JOACHIM T. HAUG AND CAROLIN HAUG Pazinato, P. G., Jauvion, C., Schweigert, G., Haug, J. T., & Haug, C. 2020: After 100 years: a detailed view of an eumalacostracan crustacean from the Upper Jurassic Solnhofen Lagerstätte with raptorial appendages unique to Euarthropoda. Lethaia, https://doi.org/10.1111/let.12382. The Solnhofen Konservat‐Lagerstätte yields a great number of remarkably preserved fossils of eumalacostracan crustaceans that help us understand the early radiation of several groups with modern representatives. One fossil from there, Francocaris grimmi Broili, 1917 is a small shrimp‐like crustacean originally described about 100 years ago as a mysidacean crustacean (opossum shrimps and relatives) from latest Kimmeridgian – early Tithonian (Upper Jurassic) of the Solnhofen Lithographic Limestones of South- ern Germany. New material with exceptionally preserved specimens, allied with mod- ern imaging techniques (mostly composite fluorescence microscopy), allows us to provide a detailed re‐description of this species. The most striking feature of Franco- caris grimmi is an extremely elongated thoracopod 7 with its distal elements forming a spiny sub‐chela. This character supports a sister group relationship of Francocaris grimmi with Eucopiidae, an ingroup of Lophogastrida, pelagic peracaridans common in marine environments throughout the world. We also discuss other supposed fossil representatives of Lophogastrida, identifying all of them as problematic at best. The structure of the sub‐chela in F. grimmi indicates an original use in raptorial behaviour. Francocaris grimmi appears to be unique in possessing such a far posterior sub‐chelate appendage as a major raptorial structure. In most representatives of Euarthropoda in which sub‐chelate appendages occur and are used for food intake, they are usually clo- ser to the mouth. □ Euarthropoda, eumalacostracan crustacean, Lophogastrida, rapto- rial, Solnhofen Lagerstätte, Upper Jurassic. Paula G. Pazinato ✉ [[email protected]], Joachim T. Haug [[email protected]], Carolin Haug [carolin.haug@palaeo-evo- devo.info], Department of Biology II, LMU Munich, Großhaderner Straße 282152, Pla- negg‐Martinsried, Germany; Clément Jauvion [[email protected]], Muséum national d'Histoire naturelle, CNRS UMR 7207, CR2P, Centre de Recherche en Paléon- tologie ‐ Paris, Sorbonne Université, 8 rue Buffon, 75005, Paris, France; Clément Jauvion [[email protected]], Muséum national d'Histoire naturelle, CNRS UMR 7590, IRD, Institut de Minéralogie, de Physique des Matériaux et de Cosmochimie, IMPMC, Sorbonne Université, 61 rue Buffon, 75005, Paris, France; Günter Schweigert [guen- [email protected]], Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1 70191, Stuttgart, Germany; Joachim T. Haug [[email protected]], Carolin Haug [[email protected]], GeoBio‐Center, LMU Munich, Richard‐Wagner‐Straße 1080333, München, Germany; manuscript received on 5/09/ 2019; manuscript accepted on 7/02/2020. Today, malacostracan crustaceans are among the sense (Barthel 1978; Barthel et al. 1990; Arratia most prominent non‐vertebrate components of fau- et al. 2015). nas, especially of the marine environments. A The lithographic limestones of Southern Germany major diversification of the group occurred in the have yielded especially numerous representatives of late Palaeozoic and many of the modern lineages Decapoda (Oppel 1862; Schweigert & Garassino have fossil representatives in the Mesozoic (Klomp- 2004; Garassino & Schweigert 2006; Schweigert 2011, maker et al. 2013). Our knowledge of the history of 2015; Schweigert et al. 2016), but also several repre- malacostracan crustaceans is, in many aspects, very sentatives of Stomatopoda (Haug et al. 2009a, 2010). detailed due to the presence of Konservat‐Lagerstät- This also includes spectacularly well‐preserved larval ten containing fossils with exceptional preservation. forms of both groups, only known from very few Among these are the Upper Jurassic lithographic other localities. Among them are modern‐looking limestones of Southern Germany, also referred to larvae of spiny lobster‐like crustaceans (Polz 1972, as Solnhofen Lithographic Limestones in a wide 1973; Haug et al. 2011a, 2014a), now extinct larval DOI 10.1111/let.12382 © 2020 The Authors. Lethaia Published by John Wiley & Sons Ltd on behalf of Lethaia Foundation This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 2 Pazinato et al. LETHAIA 10.1111/let.12382 types of spiny lobster‐like crustaceans (Polz 1996; older deposit than the overlying Zandt Member, Haug et al. 2009b, 2013; Haug & Haug 2013, 2015, according to high‐resolution biostratigraphical data 2016; 2014a), remains of larval clawed lobsters (Haug based on ammonite assemblages (Schweigert 2007). & Haug 2017), modern‐looking types of mantis The most recent discoveries of Francocaris grimmi shrimp larvae (Haug et al. 2008, 2015) and now are specimens from the Lower Tithonian Mörnsheim extinct larval forms of mantis shrimps (Haug et al. Formation of Daiting. Francocaris grimmi has not 2009a, 2015). Likewise, representatives of Peracarida yet been detected outside the Franconian Jurassic. are well known from the lithographic limestones of Southern Germany (Polz 1998, 2003, 2004). Peracarida is mainly represented by fossils of the Material and methods group Isopoda and their close relatives (Tanaidacea). A peracaridan ingroup that is more challenging con- Material cerning the fossil record is Mysidacea, which includes Stygiomysida, Mysida (opossum shrimps) All available specimens considered to represent the and Lophogastrida. The last two groups have been species Francocaris grimmi Broili, 1917 were exam- suggested as being represented in the fossil record ined. The syntype series of Broili (1917) originally (Hessler 1969; Schram 1986; Feldmann et al. 2017), comprised five specimens, of which three are still yet most of the reported fossils remain problematic. available: SNSB – BSPG 2 I 17, 1917 I 2 and 1919 I 5 This also accounts for fossils from the lithographic (Fig. 2P, C, L, respectively). The other two specimens limestones of Southern Germany. Some fossils have are lost. Broili (1917) figured three specimens (his been interpreted as representatives of Mysidacea, but fig. A: SNSB – BSPG 2 I 17, fig. B: SNSB – BSPG all of them remain questionable (Röper 2005). 1919 I 5, fig. C: specimen lost). Since the original Mysidaceans are most likely less easily preserved description, better‐preserved material became avail- as fossils, as most of them lack the well‐calcified able and is presented here. The syntype series of shields and tergites that are present in representatives Broili (1917) and specimens SNSB – BSPG 1964 of Decapoda and Isopoda. However, as the litho- XXIII 592, 1982 I 78, 1983 I 153, 1983 I 154, 1984 I, graphic limestones of Southern Germany also pre- 1986 I, 1986 I 7, 1986 I 8, 1986 I 9 are deposited in served so many ‘soft’ larval forms, we should expect the Bayerische Staatssammlung für Paläontologie the preservation of the weakly calcified mysidaceans und Geologie in Munich. Specimens GSUB A241 as well. and GSUB A242 are deposited at the Geosciences We provide here a re‐description of the rather unusual‐looking shrimp‐like crustacean Francocaris grimmi Broili, 1917 from the lithographic limestones of Southern Germany. We discuss its possible iden- tity as a mysidacean, more precisely a lophogastri- dan, and its implications for lophogastridan evolution and diversification. Geological setting The records of Francocaris grimmi come from Soln- hofen‐type Konservat‐Lagerstätten (‘Solnhofener Plattenkalke’ in German) of Franconia, Bavaria, Southern Germany (Fig. 1), a series of laminated limestones that represent deposits of carbonate parti- cles in separated basins of low energy, confined waters at the northern margin of the Tethys Sea (Robin et al. 2013; Arratia et al. 2015). The fossils originally described by Broili (1917), and most of the new material available used for this study comes from the Konservat‐Lagerstätte Zandt, east of Eich- stätt in Bavaria. Some additional material is from the ‘ ’ Fig. 1. Map of Germany, illustrating the geographical sites of latest Kimmeridgian Öchselberg limestones, near Francocaris grimmi. Occurrences of the fossils are marked with the small village of Breitenhill, which is a slightly full dots. LETHAIA 10.1111/let.12382 Eumalacostracan crustacean from Solnhofen 3 Collection of the University of Bremen. Specimens Extant specimens of Eucopia grimaldii used for SMNS 70520/1, SMNS 70520/2 and SMNS 70520/3 comparison are from the Zoologische Staatssamm- are deposited at the Staatliches Museum für Natur- lung in Munich, and the specimen of Eucopia crassi- kunde in Stuttgart, all in Germany. From the 16 anal- cornis is from the Muséum national d'Histoire ysed specimens, two are preserved in ventral naturelle in Paris. orientation (SNSB – BSPG 1964 XXIII 592 and / SMNS 70520 1) and the remaining 14 are preserved Imaging methods in lateral orientation. Most of the samples are almost complete specimens; only pieces easily lost after the The specimens were documented
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